Acta Ichthyologica et Piscatoria 51(2), 2021, 159–166 | DOI 10.3897/aiep.51.63572

Age, growth, mortality, and exploitation rate of blueline snapper, coeruleolineatus (: : ), from Dhofar Governorate, Sultanate of Oman

Dawood ALMAMARI1, Said RABIA2, Joo MYUN PARK3, Laith A. JAWAD4

1 Marine Science and Fisheries Center, Ministry of Agriculture, Fisheries Wealth and Water Resources, Muscat, Oman 2 Fisheries Research Center, Ministry of Agriculture, Fisheries Wealth and Water Resources Salalah, Salalah, Oman 3 Dokdo Research Center, East Sea Research Institute, Korea Institute of Ocean Science and Technology, Haeanro, Republic of Korea 4 Pokeno, Auckland, New Zealand http://zoobank.org/268E36E9-35EE-4A92-8DFE-1C5A16C50FC7

Corresponding author: Dawood Almamari ([email protected])

Academic editor: Adnan Tokaç ♦ Received 5 October 2020 ♦ Accepted 21 January 2021 ♦ Published 12 July 2021

Citation: Almamari D, Rabia S, Myun Park J, Jawad LA (2021) Age, growth, mortality, and exploitation rate of blueline snapper, Lutjanus coeruleolineatus (Actinopterygii: Perciformes: Lutjanidae), from Dhofar Governorate, Sultanate of Oman. Acta Ichthyologica et Piscatoria 51(2): 159–166. https://doi.org/10.3897/aiep.51.63572

Abstract

The blueline snapper, Lutjanus coeruleolineatus (Rüppell, 1838), is a significant commercial harvested from the tra- ditional fishery in the Sultanate of Oman. Deficient data on this species, however, make the management strategies challenging, especially in fisheries ecology. A total of 978 specimens were obtained from Dhofar Governorate off the coast of the Arabian Sea during the period between February 2015 and March 2016. In total, 296 sectioned otoliths were analyzed and growth was estimated from non-seasonal growth by using von Bertalanffy method. The parameters of von Bertalanffy growth function, total mortalityZ ( ), natural mortality (M), fishing mortality F( ), and exploitation rate (E) were determined and compared with those for Lutjanidae elsewhere. The age structure of male fishes was between 1 and 14 years for males while the age of females ranged from 1 to 18 years. –1 –1 Growth rate (K) was 0.21 y and 0.16 y for males and females, respectively. The hypothetical length of female (L∞ = 46 cm) was relatively higher than that of male (L∞ = 42 cm). The natural mortality (M) was 0.296, total mortality (Z) was 0.372, fishing mortality was 0.076, and exploitation rate (E) was estimated as 0.2. The presently reported study is conducted to examine this fish in relation to growth parameters by analyzing otolith structure. The results of the presently reported study will contribute towards planning the regional fishery management policies in Oman.

Keywords

Arabian Sea, blueline snapper, Dhofar, exploitation rate, mortality, otolith

Introduction ery, as well as its aquaculture in the Sultanate of Oman, raised from 211 000 t with a value of about US $ 166 mil- Fishery resources are one of the greatest marine resourc- lion in 2014 to 580 000 t with a value of about US$ 795 es in the Sultanate of Oman and have been extensively million in 2019. Oman’s coastal regions are surrounded helping fishermen for their livelihood (Al-Marzuqi 2011). by three bodies of water, including the Arabian Sea, the The total landing of artisanal, coastal, and industrial fish- Sea of Oman and, the Persian Gulf. The country is rich in

Copyright Dawood Almamari et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 160 Almamari et al.: Age, growth, mortality, and exploitation rate of Lutjanus coeruleolineatus biodiversity of marine resources and possesses a 3165 km Materials and methods long coastal line (Al-Anbouri et al. 2013). The fisheries in the Sultanate of Oman constitute a variety of commercial- Samplings and laboratory works ly important marine species that support the artisanal fish- eries sector and industrial fleet. In addition, the seafood In total, 978 specimens of blueline snapper were land- trade and related industries have investment opportunities ed by gillnets and handlines from Salalah Fish Market that help to grow and sustain the income of the fishermen (SFM) (17°01.2′N, 54°04.58′E) at Dhofar Governorate (Al-Abdessalaam 1995). from March 2015 to February 2016. Among the sam- The blueline snapper, Lutjanus coeruleolineatus (Rüp- ples collected, randomly selected 296 fish (158 females pell, 1838) (hereafter blueline snapper), is a member of and 138 males) were used for age determinations using the Lutjanidae which contains 17 genera and about sagittal otolith. A pair of otoliths (Fig. 1) were removed 113 species (Froese and Pauly 2019). There are five spe- from each specimen. The sagitta was mounted on a glass cies of the Lutjanus characterized by yellowish slide using epoxy resin (Brothers et al. 1983). The oto- body color with 4 to 8 blue longitudinal stripes (Barman lith was then grounded and Isomet low-speed diamond et al. 2014). Such snappers include Lutjanus bengalensis saw was used to make 200–300 µm thick transverse sec- (Bloch, 1790), Lutjanus kasmira (Forsskål, 1775), Lut- tions of sagittal otoliths containing the core of the otolith janus notatus (Cuvier, 1828), (Cowan et al. 1995). The sections were examined under (Bloch, 1790), and Lutjanus viridis (Valenciennes, 1846) a stereomicroscope using both reflected and transmitted (see Allen 1985). However, L. coeruleolineatus can be lights. The fish ages were determined by counting annu- distinguishable from other species of the genus by yellow li which consist of an opaque and translucent ring. The body, darker on back and whitish ventrally with distin- opaque zones were observed with reflected light while guished blue elongated stripes (7–8) on the sides, a large the translucent rings were observed in transmitted light. black spot on the , and the fish head has blue All counts were made by two readers, repeated at least spots and broken lines (Randall 1995). It is commonly two times, and then appropriate readings were selected. known as snappers and it is also called ‘Neissar’ and/or Sectioning otoliths were reading and determine the age ‘Qalaya’ in Oman. The species is properly common and based on counting the number of opaque rings along the is frequently found at a fish market and landing sites. It is dorsal edge of the sulcus acoustics (Allman et al. 2005). mainly caught with handlines, traps, and gillnets (Al-Ab- For instance, otolith with three completed annuli and a dessalaam 1995). large translucent zone will be classified as 4 age. The presently reported study was conducted to ana- lyze the stock assessment parameters of blueline snap- per, Lutjanus coeruleolineatus, using otolith micro- Estimation of growth parameters structure. Furthermore, the presently reported study attempted to determine the growth parameters based on The growth parameters of blueline snapper were estimat- males and females of blueline snapper that can reveal ed as a relation between body size and age the non-sea- signs of exploitation level. Such data clarify the state sonal von Bertalanffy growth function (Ricker 1975) us- of the stock which can be then used by decision makers ing below the formula: to manage and control the exploitation of the stock of K(t – t0) blueline snapper for sustainability over time. Hence, an Lt = L∞ × (1 – e ) essential study on the age and growth of this species is required to understand their longevity and growth rate in where Lt is length of fish at age t, L∞ is the hypothetical the Arabian Sea. length, K is the instantaneous growth coefficient and t0

Figure 1. A pair of sagittal otoliths for blueline snapper, Lutjanus coeruleolineatus. Acta Ichthyologica et Piscatoria 51(2), 2021, 159–166 161 is the hypothetical age at which length is equal to 0. The population annually through different fishing activities growth parameters of sex’s pooled, male and female data (Meyer and Schill 2014). The exploitation rate (E) were fitted using the Vonbitb program (Stamatopoulos was calculated by using Sparre and Venema (1992) and Caddy 1989) with solver parameters to reduce the equation: residual sum of squares. The longevity (life span) was estimated by (Pauly E = F × Z–1 1983) formula to estimate maximum size at age: where E is the exploitation rate, F is the fishing mortality, –1 tmax = 3K and Z is the annual instantaneous rate of mortality. where t is the longevity and K is the instantaneous growth. max Results Age validation Length frequency

In order to validate if the opaque and hyaline zone are laid Total lengths (TL) of all blueline snapper sampled ranged annually, the marginal increment analysis of the otoliths from 19.4 to 43.7 cm with a mean of 28.4 cm (±0.14). was investigated following Lessa et al. (2006): Among them, 529 female and 449 male (sex ratio of F:M = 1.18) specimens were determined, females ranged –1 MIR = (RC – RL) × (RL – RL – 1) from 20.4 to 43.7 cm TL with a mean of 29.1 cm (±0.21), which was significantly longer than the mean TL of males where MIR is the marginal increment ratio, RC is the (27.5 ± 0.16 cm, t0.05, df = 976 = −5.63, P < 0.05). otolith radius from the primordium to the edge, RL is the Analysis of the length distribution for blueline snap- otolith radius from the primordium to the last annulus, per revealed that the majority of specimens (male and fe- and RL – 1 is the otolith radius from the primordium to the male) were concentrated at length classes between 26 cm annulus next to the last one. TL and 36 cm TL. However, lengths classes less than The mean marginal increment ratio was calculated 26 cm TL and bigger than 36 cm TL were recorded by monthly and plotted to examine if the annuli of the oto- few specimens (Fig. 2). lith formed annually or not and to estimate the translucent ring growth (slow–fast) during the year.

Mortality and exploitation rate

The annual instantaneous rate of mortality (Z) was es- timated using the length converted catch curve method (Pauly 1983) and Beverton–Holt method (Beverton and Holt 1957) employed by LFDA5 program (Kirkwood et al. 2003). The fishing mortality (F) was calculated fol- lowing Pauly (1980):

F = Z − M

Natural mortality (M) was estimated from the empir- ical equation using and growth parameter K Figure 2. Length frequency distributions (cm, TL) for male (Pauly 1980) described by: and female blueline snappers, Lutjanus coeruleolineatus sam- pled from Dhofar Governorate. log M = –0.0066 – 0.279 log L∞ + 0.6543 log K + 0.4634 log T Age and growth where the T value was taken as 24°C as this value repre- sented the mean bottom water temperature in the Arabian The rings were clearly visible with a high level of read- Sea of Oman (Thangaraja 1995). The natural mortality ability (90%) and the age of the fish could be determined (M) also was measured based on longevity (Alagaraja in the majority of them. 1984; Hoenig 1984; Hewitt and Hoenig 2005) and growth The size of blueline snapper to age data were fitted for parameter K (Jensen 1996). each sex separately by using the non-seasonal von Berta- The exploitation rate can be defined as the proportion lanffy growth function (VBGF) in order to estimate the of harvestable-sized fishes that are removed from a growth pattern of this species (Fig. 3). The age structures 162 Almamari et al.: Age, growth, mortality, and exploitation rate of Lutjanus coeruleolineatus

Table 2. Age–length key of Lutjanus coeruleolineatus females from Dhofar Governorate, Sultanate of Oman.

TL Age [year] Total [cm] 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 18–20 0 20–22 3 2 5 22–24 3 17 1 21 24–26 3 15 6 24 26–28 1 5 12 4 22 28–30 6 2 13 2 23 30–32 2 3 8 5 1 1 20 32–34 5 3 2 1 11 34–36 3 2 1 2 1 9 36–38 1 2 2 1 3 1 2 12 38–40 2 1 1 1 5 40–42 1 1 1 1 1 5 42–44 1 1 Total 10 47 24 25 13 8 7 4 7 2 1 2 3 2 0 0 2 1 158 Mean 23 25 26 29 31 34 34 34 34 36 36 40 39 39 41 43 29 Figure 3. VBGF curves for male and female(n = 158) and male length (n = 138) blueline snapper, Lutjanus coeruleolineatus. [cm] estimated by otolith annuli analysis of blueline snapper Table 3. Age–length key of sex combined Lutjanus coeruleo- ranged from 1 to 14 years for males and 1 to 18 years for lineatus from Dhofar Governorate, Sultanate of Oman. females. Age distributions at each length class for male, female, and all sex combined were illustrated in Tables TL Age [year] Total [cm] 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 1, 2, and 3, respectively. Overall, the majority of the in- 18–20 0 dividuals were below age 15 (Only 3 individuals had the 20–22 4 7 11 age upper 15 years, with one specimen showing the old- 22–24 6 32 3 41 24–26 11 24 17 1 53 est age of 18 years; Table 3). The von Bertalanffy growth 26–28 1 16 18 8 2 45 function (VBGF) suggests that the male fish had a higher 28–30 8 6 21 8 1 1 45 growth rate (K = 0.21 y–1) than female. While, the hypo- 30–32 3 5 9 13 1 3 2 36 32–34 1 7 7 6 3 1 1 1 27 thetical length of the female (L∞ = 46) was significantly 34–36 0 4 2 1 2 2 1 1 1 14 higher than that of male (27.6 ± 0.33 cm, t0.05, df = 296 = 3.01, 36–38 1 2 2 2 3 1 2 13 P < 0.05). The age estimation indicates the life spans 38–40 2 1 1 1 5 (t ) are 14.2, 18.7, and 17.6 years for male, female, and 40–42 1 1 1 1 1 5 max 42–44 1 1 combined sexes, respectively. Total 22 90 50 38 32 15 13 9 7 4 2 3 4 4 0 0 2 1 296 Mean 23 24 26 28 30 33 32 33 36 35 36 36 38 36 41 43 28 length Table 1. Age–length key of Lutjanus coeruleolineatus males [cm] from Dhofar Governorate, Sultanate of Oman.

TL [cm] Age [year] Total MIR Ratios 1 2 3 4 5 6 7 8 9 10 11 12 13 14 18–20 0 Mean monthly marginal increment ratios (MIR ± SE) in 20–22 1 5 6 the otoliths of blueline snapper are shown in Fig. 4. The 22–24 3 15 2 20 24–26 8 9 11 1 29 MIR fluctuated during the year ranging between 0.48 and 26–28 11 6 4 2 23 0.79 mm and confirmed that the growth of translucent 28–30 2 4 8 6 1 1 22 ring was slow. The MIR values showed a low mean val- 30–32 1 2 1 8 1 2 1 16 32–34 1 2 4 4 2 1 1 1 16 ue (0.56) between February and July and a high mean 34–36 1 1 1 1 1 5 value (0.72) between August and December reaching a 36–38 1 1 peak value (0.79) in November. These monthly changes 38–40 0 40–42 0 in MIR over a year figured that the recognized otolith an- 42–44 0 nulus was deposited once a year. Total 12 43 26 13 19 7 6 5 0 2 1 1 1 2 138 Mean length [cm] 23 24 26 28 29 32 31 32 33 33 34 35 35 27 Mortality and exploitation rate The (VBGF) which describes male, female and com- bined sex of blueline snapper at age can be expressed as: The coefficient of total mortalityZ ( ) of blueline snapper was estimated by the length converted catch curve meth- –0.21(t + 0.22) 2 Male: TL [cm] = 42 [1 – e ], (r = 0.70) od (Z = 0.393) for combined sex (Fig. 5). The Z value estimated through the Beverton-Holt method revealed as –0.16(t + 0.15) 2 Female: TL [cm] = 46 [1 – e ], (r = 0.78) Z = 0.350. Therefore, the mean value of Z value from two methods was taken and hence the Z = 0.372 was used for –0.17(t + 0.18) 2 Combined sex: TL [cm] = 44 [1 – e ] (r = 0.73) the calculation of fishing mortality. The natural mortality Acta Ichthyologica et Piscatoria 51(2), 2021, 159–166 163

Figure 5. Length converted catch curve of blueline snapper, Lutjanus coeruleolineatus. Figure 4. Monthly variations in the mean (±SE) marginal oto- lith increment (MIR) [mm] of blueline snappers Lutjanus coe- single annual ring formation (Cappo et al. 2000; Newman ruleolineatus in the Dhofar Governorate coast of the Arabian et al. 2000a, b; Burton 2002), corresponding to the major- Sea from March 2015 to February 2016. ity of tropical fishes (e.g., Pilling et al. 2000). In addition, many previous studies have considered Lutjanidae fishes (M) calculated from the empirical equation using tem- growing faster especially during their first year of their perature and growth parameter (K), longevity, and growth life history (Newman et al. 1996; Newman et al. 2000a; parameter (K) was 0.47, 0.26, 0.24, 0.25, and 0.26 respec- Kritzer 2004; Grandcourt et al. 2006). tively. Thus, the mean values of natural mortality (M) Age–length frequency distributions for the presently from all methods were taken and equal to 0.296. There- reported blueline snapper revealed that only 1% of the fore, fishing mortality (F) was equal to 0.076 by subtract- samples were older than age 15. Although the sample ing the natural mortality (M) from the total mortality (Z). size in this study was not enough for validating popu- The exploitation rate (E) is estimated as 0.2. lation-level age structure. Male’s age ranged from 1 to 14 years, while females were between 1 and 18 years showing toward higher age distribution. The age struc- Discussion tures were variable between sexes, with the female hav- ing the highest longevity at 18.7 years. This estimation The basic biological studies such as morphometric, age of longevity is similar to those attained by Lutjanus car- and growth parameters, and reproductive characteristics ponotatus (Richardson, 1842) sampled within the Great are important in understanding the life history of the fish- Barrier (Newman et al. 2000b; Kritzer 2004). On es. The maximum fish length of the presently reported other hand, males blueline snapper had less life span species was 43.7 cm TL while previous estimation (Ran- (14.2 years) than females (18.7 years). Such results were dall 1995) yielded 40 cm TL. We found a significant dif- also reported by Heupel et al. (2009) with male Lutja- ference in the mean total length between males and fe- nus gibbus (Forsskål, 1775), Aprion virescens Valenci- males, with females having bigger body sizes than males. ennes, 1830, and Lutjanus fulvifamma displaying short- This observation coincides with other Lutjanidae fishes er life spans of maximum ages of 12, 16, and 17 years, including Lutjanus fulviflamma (Forsskål, 1775) from respectively. Shorter longevities in male snappers were southern Arabian Gulf (Grandcourt et al. 2006) and Mafia also recorded in (Quoy et Gaimard, 1824) Island in Tanzania (Kamukuru et al. 2005), and Lutjanus (see Newman et al. 2000b), Lutjanus guttatus (Forsskål, campechanus (Poey, 1860) from the Atlantic waters of 1775) (see Amezcua et al. 2006), and L. fulvifamma (see the southeastern USA (White and Palmer 2004). These Grandcourt et al. 2006). results may be related to differences in be- The asymptotic length (L∞) of blueline snapper esti- tween sexes, such as differences in the level of surplus mated was 44 cm TL for combined sex. The growth rate energy between reproduction and somatic growth (Rljns- of males (K = 0.21) was higher than females (K = 0.16). dorp and Ibelings 1989). Such finding is relatively in agreement with other- Lut In the presently reported study, sectioned otoliths janidae fishes (Newman et al. 2000b; Kamukuru et al. were used to estimate growth parameters. Various meth- 2005; Heupel et al. 2010; Shimose and Nanami 2014). ods have been used for estimating growth parameters of Additionally, Newman et al. (2000b) have clarified that Lutjanus species, but reading otolith sections are one of the variations in the growth rate of L. carponotatus were the best application for annuli determination due to its ac- probably linked to gonad maturation and sexual ma- curacy and reliability (Newman et al. 1996). Many previ- turity. Moreover, the growth rate observed for blueline ous listings suggested that snapper species generally have snapper was rapid during 3–4 years of age attaining over 164 Almamari et al.: Age, growth, mortality, and exploitation rate of Lutjanus coeruleolineatus

25.0 cm TL for combined sexes and then became slower. The presently reported study estimation of the annual Similar to this result, a growth study of blacktail snap- instantaneous rate of mortality (Z) for the blueline snap- per, (Forster, 1801), from the Yaeyama per in the Dhofar Governorate was 0.372 and the fishing Islands, Okinawa, Japan revealed that such rapid growth mortality (F) was 0.076. Total mortality calculations is probably due to their maturation (Shimose and Nanami based on the length converted catch curve method has 2014). Shimose and Tachihara (2005) also suggested that been recorded high value (0.393) than the Beverton– reproductive status is often considered to be the main rea- Holt method (0.350). Estimates were greatly lower than son for inhibiting the growth of small Lutjanus species. that found by Burton (2002) (Z = 0.49) and higher than Some Lutjanus snappers have a relatively long life Mason and Manooch (1985) (Z = 0.33) for L. analis. The span and larger maximum size (≥50.0 cm TL), often ex- optimum exploitation rate (E) is assumed to be close ceed 40 years such as (Forsskål, 1775) to 0.5 (Gulland 1971). We compared the exploitation nearly close to 56 years (Marriott and Mapstone 2006), status of blueline snapper (presently-reported study) L. campechanus to 45 years (White and Palmer 2004), with results from previous studies on different families (Bloch et Schneider, 1801) to 48 for demersal fish in Oman. The exploitation rate E( ) of years, Bloch, 1790 to 42 years blueline snapper was 0.2 showing the stock at a lower (Fry et al. 2009), and Lutjanus fulvifamma (Forsskål, optimum level of exploitation. Similar to these results, 1775) to 40 years (Shimose and Tachihara 2005). In con- Al-Mamry et al. (2011) stated that the exploitation sta- trast, some smaller Lutjanus snappers had a short life tus of sea bream species, spinifer (Forsskål, span; for example, the age of Lutjanus ehrenbergii (Pe- 1775) was optimal and the stock was in a healthy sta- ters, 1869) reaches to around 12 years (Grandcourt et al. tus. On the contrary, Abd El Barr (2016) specified that 2011), Lutjanus synagris (Linnaeus, 1758) to 19 years overexploitation of emperors, groupers, sea bream, and (Luckhurst et al. 2000), (Castelnau, 1873) rabbit fish due to the rise number of boats and increase to 12 years (Newman et al. 1996), and fishing activities on those species. Few studies on ex- (Cuvier, 1816) to 22 years (Newman et al. 2000a). ploitation status of demersal fish caught from the- ar Estimates of natural mortality (M) for the presently tisanal fishery in Oman. Therefore, further studies are reported species was 0.296 which is similar to those for required to compare the exploitation status between de- other Lutjanidae fishes. Generally, manyLutjanus species mersal species. are long-lived fishes and have a low value of the instanta- The study was competent to investigate age structure, neous rate of natural mortality; for instance, natural mor- growth rate, and the mortality of blueline snapper, L. coe- tality of Lutjanus analis ranged from 0.28 and 0.49 (Bur- ruleolineatus in Omani waters (Arabian Sea). The length ton 2002) and L. campechanus was 0.11 (Topping and and distributions between females and males were Szedlmayer 2013). Newman et al. (2000b) also investi- comparable to those reported in L. fulvifamma from gated the life histories of two species (L. carponotatus Southern Arabian Gulf, Mafia Island in Tanzania, and and L. vitta) and found the natural mortalities of those Lutjanus campechanus from the northern Gulf of Mexico species were between 0.20–0.21 and between 0.34–0.35, suggesting females having bigger and heavier body than respectively. Moreover, the natural mortality of L. griseus males (Wilson and Nieland 2001; Kamukuru and Mga- was between 0.15 and 0.50 (Fischer et al. 2005), ya 2004). Estimation of growth rate was rapid during the L. fulvifamma was around 0.29 (Grandcourt et al. 2006) first 3–4 years of age attaining over 25.0 cm TL and then and L. carponotatus was between 0.18 and 0.30 (Heupel became slower. This study provides basic biological data et al. 2010). While, Pauly (1980) estimated the natural that is useful in sustainable fishery management of this mortality of fishes via Von Bertalanffy growth parameters species in the Sultanate of Oman.

(L∞ or W∞, and K) and mean environmental temperature (T), and assumed that there is a relation between fish size and natural mortality. Environmental factors such as sea- Acknowledgments water temperature possibly affect life; e.g., warm water will have higher mortality rates than an equivalent My great thank to the Ministry of Agriculture, Fisheries animal living in cooler water (Pauly 1980). Whereas es- Wealth and Water Resources, Sultanate of Oman for pro- timation of natural mortality for the presently reported viding me all facilities to make this research. My kind species that lives in warm waters have a similar range of appreciation goes to my colleagues at Marine Science and that species that inhabiting colder waters. Fisheries Center for their recommendations.

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